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An Ecological Comparison of Small Mammal Communities in California

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An Ecological Comparison of Small Mammal Communities in California An Ecological Comparison of Small Mammal Abstract: Our studies in similar scrub habitats Communities in California and Chile1 in California and Chile reveal some interesting differences between these two regions in the structure of their small mammal communities. The Chilean fauna is less diverse, with fewer species 2 William E. Glanz and Peter L. Meserve per site, and possibly more extreme density fluc- tuations. Chilean rodents are more strongly associated with areas of high shrub and rock cover, while California species show a greater variety of habitat preferences. Chile has more insectivorous species, and California has more seed-eating specialists. Some of these differ- ences may be related to biogeographic and climatic factors, while others may reflect a longer history of human disturbance in Chile. Similar climates often seem to favor the evo- FIELD SITES AND METHODS lution of similar organisms, and the Mediterranean -climate regions of the world have provided numer- We have ecological data from a variety of ous examples of such evolutionary convergence. communities in each region, but will restrict our Of the five regions with this climate, California discussion here to results from three vegetational and Chile have been compared very extensively types which we have studied intensively: dry (Mooney 1977), particularly in terms of the mor- coastal scrub, moist coastal scrub, and evergreen phological, physiological, and ecological attri- chaparral. For each of these, the vegetation butes of their dominant organisms. Striking structure and life forms at our sites were closely similarities between these two regions have been matched between continents. found in their vegetation structure and community patterns (Mooney and Dunn 1970; Parsons and The dry coastal scrub localities were studied Moldenke 1975; Parsons 1976), their birds (Cody by Meserve near Irvine, Orange County, California 1973 and 1974), and their lizards (Fuentes 1976). in 1970-71 and at Fray Jorge National Park, Coquimbo Province, Chile in 1973-74. Detailed In a variety of projects during the past decade descriptions were provided by M'Closkey (1972) (Meserve 1972, 1976a, 1976b, 1981a, 1981b; Glanz and Meserve (1972 and 1976a) for the California 1977a and 1977b; Meserve and Glanz 1978), we have site, and by Fulk (1975) and Meserve (1981a) for studied certain communities of small mammals in the Chilean site. Both study areas were at about the Mediterranean zones of California and Chile. 200 m elevation, within 5 km of the coast, and In this paper we will summarize our results by were located on sandy, well-drained soils. comparing our three best-documented sites on each Although mean rainfall at the California site continent. We will outline some aspects of these (about 300 mm) was considerably greater than at communities that suggest convergence in morphology the Chilean site (127 mm), their similar topo- and ecology. Our emphasis, however, will be on graphy, seasonality, and soils apparently pro- the ecological differences that we have found duced remarkably convergent vegetation, with between the two regions, especially those relating approximately 60 percent shrub cover dominated by to community diversity, habitat use, and diet. drought-deciduous species. These dissimilarities suggest some major differ- ences in the functional roles of mammals in these The moist coastal sites studied by Glanz communities, some contrasting responses of these (1977a and 1977b) were at Camp Pendleton Marine animals to vegetation structure, natural preda- Base, San Diego County, California, and at tors, and human disturbance, and some potentially Zapallar and Los Molles (two sites), Aconcagua important differences in the evolutionary his- Province, Chile. Although the rainfall at these tories of the two regions. We hope that our dis- localities (225 to 350 mm) was comparable to that cussion of these topics will stimulate further of the California dry coastal site, they were on research on these ecosystems and their small slopes more directly facing the ocean and received mammal faunas. more coastal fog. This greater moisture plus the higher clay content of the soils resulted in greater average shrub cover (74 percent), also 1 Presented at the Symposium on Dynamics and dominated by drought-deciduous species, but Management of Mediterranean-type Ecosystems, including more evergreen shrubs than the dry June 22-26, 1981, San Diego, California. coastal sites. All study areas were between 20 and 60 m elevation and within 1 km of the coast. 2 Assistant Professor of Zoology, University of Maine, Orono, Maine 04469; and Assistant Pro- The evergreen chaparral sites sampled by fessor of Biological Sciences, Northern Illinois Glanz (1977a and 1977b) were at Echo Valley, University, DeKalb, Illinois 60115. near Descanso, San Diego County, California, and Gen. Tech. Rep. PSW-58. Berkeley, CA: Pacific Southwest Forest and Range 220 Experiment Station, Forest Service, U.S. Department of Agriculture; 1982. at Fundo Santa Laura, near Tiltil, Santiago Table 1. Occurrence of each small mammal species Province, Chile. Both were at 900 to 1000 m ele- at the California (above) and Chilean (below) vation, had 550 to 600 mm mean annual rainfall, sites. Sites are dry coastal (DC), moist coastal and were dominated by evergreen sclerophyllous (MC), and evergreen chaparral (EC). Abundance shrubs. Plots trapped at these sites averaged 75 values are: X = present, but uncommon; L = locally percent shrub cover. Many additional aspects of common; C = common or abundant. Species Codes are the evergreen chaparral and moist coastal sites used in figures 1 and 2. were studied during the Mediterranean Scrub Project of the International Biological Program, and are Community Type described in detail in Thrower and Bradbury (1977). Species (and Code) DC MC EC CALIFORNIA Our field techniques were designed to sample most mammal species up to 500 gm body size in each INSECTIVORA community. Populations were live-trapped on large Soricidae (1.4 to 2.2 ha) grids at all sites except the Notiosorex crawfordi X X Chilean moist coastal localities, and snap-trapped RODENTIA on small (usually 0.4 ha) grids at all sites except Sciuridae to California dry coastal area. Population den- Eutamias merriami X sities were determined by mark-recapture tech- Spermophilus beecheyi L L niques, and by correlating snap-trap success with Geomyidae live-trapping results. A variety of physical and Thomomys bottae X X vegetational measurements were used to character- Heteromyidae ize the habitat preferences of each species. Dipodomys agilis (Dpa) C C Meserve (1976b, 1981b) categorized each trap Perognathus californicus (Pgc) X X station according to its shrub, herb, and bare Perognathus fallax (Pgf) X C ground cover values, and then conducted 2 x k association analyses (Simpson, Roe, and Lewontin Perognathus longimembris (Pgl) C 1960) between presence/absence data of each Cricetidae species at trap stations and categories of vege- Microtus californicus (Mcc) X C X tation cover. Glanz measured 30 habitat variables Neotoma fuscipes (Ntf) L C C at each trap station, scored each mammal capture Neotoma lepida (Ntl) X C X for the characteristics of that trap station, and Peromyscus boylii (Pmb) L then compared the habitat measures of stations Peromyscus californicus (Pmc) C C C selected by each mammal species with those of all Peromyscus eremicus (Pme) C C X stations trapped at each site. To assess the Peromyscus maniculatus (Pmm) C C C degree of habitat differentiation among species Reithrodontomys megalotis (Rtm) C X X at each site, Glanz also conducted discriminant Muridae analyses using capture records and programs in Mus musculus X X the SPSS computer library. Total Species 12 13 13 Meserve compared arboreal vs. terrestrial Common Species 7 8 6 habitat utilization at the dry coastal sites CHILE using smoked cards placed on the grant and at 25, 50, 100 and 200 cm above ground in the most common MARSUPIALIA shrub species. Tracks on the cards were identi- Didelphidae fied to species and individual using foot charac- Marmosa elegans (Mae) X C X teristics and toe-clip marks (Meserve 1976b and RODENTIA 1981b). Abrocomidae Abrocoma bennetti (Abb) X C Food habits were assessed by microscopic Octodontidae examination of stomach contents and fecal pellets Octodon degus (Ocd) C X C at most sites, and by fecal analysis only at the Octodon lunatus (Ocl) L L California dry coastal site. Meserve (1976a and Spalacopus cyanus L X 1981a) homogenized the sample first, subsampled Cricetidae and boiled the material in Hertwig's solution Akodon longipilis (Akl) C C C (Baumgartner and Martin 1939), and then identi- Akodon olivaceus (Ako) C C X fied cell fragments under a microscope at 100X. Oryzomys longicaudatus (Orl) X X X Glanz (1977) quantified stomach content fractions Phyllotis darwini (Phd) C X X first at 40X, subsampled each fraction, and then followed the above procedure. These variations Muridae in methods may limit the precision of our compar- Rattus rattus (Rra) L isons, but should not alter the general conclu- Total Species 7 8 10 sions we present in this paper. Common Species 4 5 5 FAUNAL COMPOSITION AND DIVERSITY each community are listed in table 1. Several points are obvious from this list. First, both The species of small mammal recorded from faunas are dominated by rodents; the only excep- 221 tions are one shrew, Notiosorex crawfordi, in the two continents (p>.10; U-test). The Chilean California and one mouse opossum, Marmosa elegans site had more extreme variations (coefficient of in Chile. Next, the two faunas are phylogeneti- variation = 48.9 in Chile, 16.8 in California), cally distinct. Among the rodents, only one with Akodon olivaceus declining from high densi- native family (Cricetidae) is shared between the ties (32/ha) early in the study and Octodon degus two regions, although each also contains species reaching 120/ha near the end (Meserve 1981b). of murid rodents introduced by humans from Fulk (1975) live-trapped the Fray Jorge site in Eurasia.
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